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''Mimodactylus'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of mimodactylid
pterodactyloid Pterodactyloidea (derived from the Greek words ''πτερόν'' (''pterón'', for usual ''ptéryx'') "wing", and ''δάκτυλος'' (''dáktylos'') "finger" meaning "winged finger", "wing-finger" or "finger-wing") is one of the two traditional ...
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 to ...
from the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
of what is now Lebanon.


Discovery and naming

In the ''
Lagerstätte A Lagerstätte (, from ''Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that exhibits extraordinary fossils with exceptional preservation—sometimes including preserved soft tissues. These for ...
'' of
Hjoula Hjoula ( ar, حجولا) is a municipality in the Byblos District of Keserwan-Jbeil Governorate, Lebanon. It is 70 kilometers north of Beirut. Hjoula has an elevation of between 920 and 1,100 meters above sea level. Hjoula has a total land area o ...
, ten kilometres inland from the port of
Byblos Byblos ( ; gr, Βύβλος), also known as Jbeil or Jubayl ( ar, جُبَيْل, Jubayl, locally ; phn, 𐤂𐤁𐤋, , probably ), is a city in the Keserwan-Jbeil Governorate of Lebanon. It is believed to have been first occupied between 880 ...
, commercial fossil hunters regularly make important discoveries. A rare pterosaur skeleton was acquired by an anonymous benefactor and donated to the ''Mineral Museum'' at
Beirut Beirut, french: Beyrouth is the capital and largest city of Lebanon. , Greater Beirut has a population of 2.5 million, which makes it the third-largest city in the Levant region. The city is situated on a peninsula at the midpoint o ...
, part of the ''
Université Saint-Joseph Saint Joseph University of Beirut ( French: ''Université Saint-Joseph de Beyrouth'', abbreviated to and commonly known as "USJ") is a private Catholic research university located in Beirut, Lebanon, which was founded in 1875 by French Jesuit mis ...
''. It was prepared in Canada by Luke Allan Lindoe, a technician of the
University of Alberta The University of Alberta, also known as U of A or UAlberta, is a public research university located in Edmonton, Alberta, Canada. It was founded in 1908 by Alexander Cameron Rutherford,"A Gentleman of Strathcona – Alexander Cameron Rutherfor ...
. Since 2017, the museum displays two life-size models of the animal, one standing, the other flying, and a
hologram Holography is a technique that enables a wavefront to be recorded and later re-constructed. Holography is best known as a method of generating real three-dimensional images, but it also has a wide range of other Holography#Applications, applic ...
. In 2019, the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
''Mimodactylus libanensis'' was named and described by Alexander Wilhelm Armin Kellner, Michael Wayne Caldwell, Borja Holgado,
Fabio Marco Dalla Vecchia Fabio is a given name descended from Latin ''Fabius'' and very popular in Italy and Latin America (due to Italian migration). Its English equivalent is Fabian. The name is written without an accent in Italian and Spanish, but is usually accented ...
, Roy Nohra, Juliana Manso Sayão and
Philip John Currie Philip John Currie (born March 13, 1949) is a Canadian palaeontologist and museum curator who helped found the Royal Tyrrell Museum of Palaeontology in Drumheller, Alberta and is now a professor at the University of Alberta in Edmonton. In the ...
. The generic name combines a reference to the MIM, the usual acronym of the museum, with a Greek δάκτυλος, ''daktylos'', "finger". The name was intended to honour the philanthropist as well. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
refers to the provenance from
Lebanon Lebanon ( , ar, لُبْنَان, translit=lubnān, ), officially the Republic of Lebanon () or the Lebanese Republic, is a country in Western Asia. It is located between Syria to the north and east and Israel to the south, while Cyprus li ...
. The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
, MIM F1, was excavated from a marine deposit of the Sannine Chalk dating from the late
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
, about ninety-five million years old. The layers were deposited off the coast of the Afro-Arabic plate, in the south of the
Tethys Sea The Tethys Ocean ( el, Τηθύς ''Tēthús''), also called the Tethys Sea or the Neo-Tethys, was a prehistoric ocean that covered most of the Earth during much of the Mesozoic Era and early Cenozoic Era, located between the ancient continents ...
. It consists of a relatively complete skeleton with skull and lower jaws. It lacks some vertebrae and some pelvic elements. The skeleton is largely articulated and partly three-dimensionally preserved but the rear of the skull is strongly compressed. It is a juvenile individual. It represents the most complete pterosaur skeleton ever found on the Afro-Arabic plate. The University of Alberta made a cast, with inventory number MN 7216-V.


Description


Size and distinguishing traits

The holotype of ''Mimodactylus'' has a wingspan of 132 centimetres but the exemplar was far from being full-grown. The describing authors identified some distinguishing traits. Two of these are
autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to t ...
, unique derived characters. The humerus has a rectangular deltopectoral crest. The humerus has less than half of the length of the second phalanx of the fourth (wing) finger. Additionally the fossils shows a combination of traits that is unique for the
Ornithocheiroidea Ornithocheiroidea (or ornithocheiroids) is a group of pterosaurs within the extinct suborder Pterodactyloidea. They were typically large pterosaurs that lived from the Early to Late Cretaceous periods (Valanginian to Maastrichtian stages), with f ...
. The palate has a distinct ridge. The upper jaws each have eleven teeth and the lower jaws each ten. The shoulderblade is somewhat longer than the
coracoid A coracoid (from Greek κόραξ, ''koraks'', raven) is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is prese ...
. The humerus is 30% longer than the thighbone. The deltopectoral crest covers 40% of the shaft length of the humerus. From the ''Lagerstätte'' of Hjoûla two other pterosaur specimens are known. One of these is the holotype of ''
Microtuban ''Microtuban'' is an extinct genus of azhdarchoid pterosaur known from the Late Cretaceous of northern Lebanon. Discovery ''Microtuban'' is known only from a partially preserved skeleton lacking the skull, holotype SMNK PAL 6595. It was acquire ...
'', the other is specimen MSNM V 3881. According to the describing authors, both clearly differed in proportions from the ''Mimodactylus'' holotype, proving that they represented different taxa.


Skeleton


Skull

The snout of ''Mimodactylus'' is wide but not rounded in front, ending in a pointed tip. The upper jaw bears eleven teeth. These are somewhat transversely flattened, but not dagger-shaped or serrated. They are conical with an oval cros-section. Their outer side is convex. Sharp cutting-edges or ''carinae'' are lacking. Their apices are sharp and slightly bent inwards. They possess a cingulum, a thickened ridge at the crown base. The first tooth is smaller, with a circular cross-section. Subsequent teeth are longer, straight and widely spaced. They are only present in the front half of the jaw. The palate forms a concave plate with a small ridge. The ''choanae'', the internal nostrils, are large and separated by the vomers. The ''fenestrae postpalatinae'' are elongated and egg-shaped in profile, as with ''
Hongshanopterus ''Hongshanopterus'' is a genus of pterodactyloid pterosaur from the Lower Cretaceous Jiufotang Formation of Liaoning, China. The type species ''Hongshanopterus lacustris'' was in 2008 named and described by Wang Xiaolin, Diogenes de Almeida C ...
''. The
quadrate Quadrate may refer to: * Quadrate bone * Quadrate (heraldry) * Quadrate lobe of liver * Quadrate tubercle The quadrate tubercle is a small tubercle found upon the upper part of the femur. It serves as a point of insertion of the quadratus femori ...
is angled at 150° relative to the jaw edge. The lower jaw has a preserved length of 105 millimetres. It bears ten teeth for a total of forty-two in the head as a whole. The lower jaws have a central forwards pointing odontoid process in front. Of the
hyoid bone The hyoid bone (lingual bone or tongue-bone) () is a horseshoe-shaped bone situated in the anterior midline of the neck between the chin and the thyroid cartilage. At rest, it lies between the base of the mandible and the third cervical vertebr ...
two first ceratobranchialia have been preserved, thin elongated fork-shaped elements.


Postcrania

The front vertebrae of the back are not fused into a
notarium Notarium or os dorsale is a bone consisting of the fused vertebra of the shoulder in birds and some pterosaurs. The structure helps brace the chest against the forces generated by the wings. In birds, the vertebrae are only in contact with adjace ...
. A series of seven tail vertebrae is visible. These quickly diminish in size towards the rear, indicating that the tail was very short. The breastbone is rounded in front. Its cristospina, central crest on the underside, is short and deep. The shoulder blade is robust and constricted. It is longer than the
coracoid A coracoid (from Greek κόραξ, ''koraks'', raven) is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is prese ...
, different from the condition in the
Istiodactylidae Istiodactylidae is a small family of pterosaurs. This family was named in 2001 after the type genus ''Istiodactylus'' was discovered not to be a member of the genus ''Ornithodesmus''. Systematics and distribution Remains of taxa that can be co ...
. The coracoid facet touching the breastbone is concave, with a process extending to behind, again different from the isiodactylid morphology. The wings are conspicuously elongated, with a low aspect ratio. The humerus is fifty-two millimetres long. Its deltopectoral crest has distinctive straight distal edge. This crest covering 40% of shaft length exceeds the ratio in all other Ornithocheiroidea, except the
Pteranodontidae The Pteranodontidae are a family of large pterosaurs of the Cretaceous Period of North America and Africa. The family was named in 1876 by Othniel Charles Marsh. Pteranodontids had a distinctive, elongated crest jutting from the rear of the head ( ...
. The pteroid, the bone supporting the forward wing membrane or propatagium, is very long, somewhat longer even than the humerus. It clearly points towards the neck and articulates with the proximal syncarpal; this had been a point of contention among researchers. The first and second phalanx of the fourth finger are relatively longer than with istiodactylids. The fourth phalanx is curved to behind.


Phylogeny

In 2019, ''Mimodactylus'' was, within the
Lanceodontia Ornithocheiromorpha (from Ancient Greek, meaning "bird hand form") is a group of pterosaurs within the suborder Pterodactyloidea. Fossil remains of this group date back from the Early to Late Cretaceous periods (Valanginian to Turonian stag ...
, placed in the
Istiodactyliformes Ornithocheiromorpha (from Ancient Greek, meaning "bird hand form") is a group of pterosaurs within the suborder Pterodactyloidea Pterodactyloidea (derived from the Greek words ''πτερόν'' (''pterón'', for usual ''ptéryx'') "wing", a ...
. More precisely, it would be a member of the
Mimodactylidae Mimodactylidae is a group or family of pterosaurs known from Cretaceous deposits in East Asia and the Middle East. They are phylogenetically more closely related to the Istiodactylidae than to the Ornithocheiridae or the Anhangueridae, forming th ...
, as
sister species In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
of ''
Haopterus ''Haopterus'' is a genus of pterodactyloid pterosaur from the Barremian-Aptian-age Lower Cretaceous Yixian Formation of Liaoning, China. Its fossil remains dated back 124.6 million years ago. Discovery and naming It was in 2001 named by Wang Xia ...
''. The cladogram below is a topology recovered by Kellner ''et al.'' (2019). In the analyses, they recovered ''Mimodactylus'' as the sister taxon of ''
Haopterus ''Haopterus'' is a genus of pterodactyloid pterosaur from the Barremian-Aptian-age Lower Cretaceous Yixian Formation of Liaoning, China. Its fossil remains dated back 124.6 million years ago. Discovery and naming It was in 2001 named by Wang Xia ...
'' within the family
Mimodactylidae Mimodactylidae is a group or family of pterosaurs known from Cretaceous deposits in East Asia and the Middle East. They are phylogenetically more closely related to the Istiodactylidae than to the Ornithocheiridae or the Anhangueridae, forming th ...
, and placed within the more inclusive group
Istiodactyliformes Ornithocheiromorpha (from Ancient Greek, meaning "bird hand form") is a group of pterosaurs within the suborder Pterodactyloidea Pterodactyloidea (derived from the Greek words ''πτερόν'' (''pterón'', for usual ''ptéryx'') "wing", a ...
.


Paleobiology

The
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
of ''Mimodactylus'' consisted of the islands and archipelagos situated on the wide chalk plateau extending from the north coast of the Afro-Arabische plate, in the
Neotethys The Tethys Ocean ( el, Τηθύς ''Tēthús''), also called the Tethys Sea or the Neo-Tethys, was a prehistoric ocean that covered most of the Earth during much of the Mesozoic Era and early Cenozoic Era, located between the ancient continents ...
. The dentition of ''Mimodactylus'' differs from all other known pterosaurs. Its short unserrated straight conical teeth would have been useful to crack the exoskeletons of
arthropods Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
. That could indicate it was an insectivore, but insects are largely absent from the ''Lagerstätte'' layers. A lack of plant fossils proves that the location was far from land. Its elongated wings did not provide the manoeuvrability needed to catch fast flying insects. However, they would allow for a stable flight during dynamic soaring over the sea surface. The site is rich in fossils of
crustaceans Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
belonging to the
Decapoda The Decapoda or decapods (literally "ten-footed") are an order of crustaceans within the class Malacostraca, including many familiar groups, such as crabs, lobsters, crayfish, shrimp and prawns. Most decapods are scavengers. The order is esti ...
. Such shrimp-like creatures could have been scooped from the water surface by the wide beak, similar to the way some extant ducks, herons and
shoebill The shoebill (''Balaeniceps rex'') also known as the whalebill, whale-headed stork or shoe-billed stork, is a very large long-legged wading bird. It derives its name from its enormous shoe-shaped bill. It has a somewhat stork-like overall form ...
s catch prey.


See also

*
Timeline of pterosaur research This timeline of pterosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and taxonomic revisions of pterosaurs, the famed flying reptiles of the Mesozoic era. Although pterosaurs w ...


References

{{Portal bar, Paleontology, Cretaceous, Lebanon Pteranodontoids Fossil taxa described in 2019